侵染中国天南星科药用植物马铃薯Y病毒属成员的分子生物学研究
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摘要
天南星科植物中有许多种类是中国传统的药用植物。几千年来，这些药用植物都是野生的，但是随着人们对这些药用植物需求的增加，逐渐开始人工种植，而病毒病害是这些植物人工种植的一个严重问题。最近我们鉴定了一种侵染半夏(Pinellia ternata(Thunb．)Ten．ex Breitenb．)马铃薯Y病毒属(genus Potyvirus)成员，序列分析表明这种病毒与大豆花叶病毒(Soyben mosaic virus，SMV)十分相似，于是我们建议将这种病毒鉴定为SMV半夏分离物，并命名为SMV-P。
为了进一步鉴定侵染天南星科药用植物的Potyvirus属成员，我们首先用RT-PCR的方法对来自全国不同地区样品进行了检测。在供检的20个样品中，我们从7个样品中检测到了3种Potyvirus属病毒，其中5个半夏病样中含有SMV-P，1个来自江苏射阳的半夏病样含有SMV-P和芋花叶病毒(Dasheen mosaic virus，DsMV)，该分离物暂命名芋花叶病毒射阳分离物(DsMV-SY1)，另一个来自广西南宁水半夏(Tyonium flagellifrme)病样含有魔芋花叶病毒(Konjak mosaic virus，KoMV)，该分离物暂命名为魔芋花叶病毒水半夏分离物(KoMV-T)。这是KoMV侵染我国天南星科植物的首次报道。
我们测定了大豆花叶病毒杭州半夏分离物3(SMV-PHZ3)单链正义RNA基因组全序列，其全基因组由9734个核苷酸组成(不包括3’-polyA)，编码一个山3105个氨基酸组成的多聚蛋白，分子量为353.8kDa。在基因组编码的大部分蛋白中，SMV-PHZ3与大豆花叶病毒杭州半夏分离物1(SMV-PHZ1)和大豆花叶病毒杭州大豆分离物(SMV-GHZ)氨基酸序列相似性约为90％，而在P1蛋白部分，SMV-PHZ3和DsMV的氨基酸序列相似性最高。结果进一步证实，SMV-PHZ3和SMV应该是同种病毒不同分离物。同时还测定了其余5种SMV-P分离物3’术端序列，并在此基础上研究了病毒不同分离物的分子差异和进化关系。
我们测定了侵染射阳半夏的DsMV-SY1 3’末端1743个核苷酸。系统进化树进行分析表明DsMV-SY1，香果兰花叶病毒(Vanilla mosaic viurs，VanMV)和其它DsMV亲缘关系较高，成簇明显，而且DsMV分离物之间的关系和SMV分离物之间的关系很相似。Western blot分析表明SMV-P和DsMV有较弱的血清学反应。
我们测定了KoMV-T基因组全序列，其全基因组由9596个核昔酸组成，3’-末端具有poly(A)尾。KoMV-T和另一分离物F的P3蛋白氨基酸序列比较发现，两个分离物P3基因其余部分相似性比较高，但是在基因的N端度了中间部分KoMV-T比KoMV-F P3多了6个氨基酸，这可能与致病性有关。CP蛋白氨基酸序列系统进化树分析显示，KoMV分离物以及马蹄莲花叶病毒(Zantedeschia mosaic virus，ZaMV)分离物和JHMV分离物形成远源相关，与其它病毒无明显进化相关性。
用GST-P1抗血清免疫金标记发现SMV-P P1蛋白定位在细胞质和叶绿体，而在线粒体、细胞核、液泡、细胞壁等均没有观察到金颗粒标记，在阴性对照中几乎没有观察到金颗粒。P1蛋白进入叶绿体可能也是SMVP致病的一种方式。我们用酵母双杂交的方法验证P1蛋白与SMV-P编码的蛋白不存在互作。进而以SMV-P P1蛋白为诱饵从半夏酵母双杂交cDNA文库中筛选到一个候选寄主蛋白。通过质粒回收后，序列分析表明插入的部分基因与光合作用电子传递链细胞色素b_6／f复合体中Rieske Fe／S蛋白氨基酸同源性最高。用5’-RACE的方法获得该基因的全长序列。体外免疫共沉淀进一步证明P1和Rieske Fe／S蛋白互作。通过缺失突变分析表明P1-Fe／S互作的区域主要集中在SMV-P P1 N端，包括1—82位氨基酸，Rieske Fe／S蛋白的缺失突变表明Rieske Fe／S蛋白的信号肽也参与了与P1 N端34—82位氨基酸部分的互作。在大豆和马蹄莲中也同样存在SMV-P1-SFe／S和DsMV-P1-CFe／S的相互作用。共聚焦扫描显微镜下观察到GFP-P1融合蛋白在转基因拟南芥的细胞质中表达，但P1转基因拟南芥没有产生明显症状。
为了进一步研究SMV-P各基因功能，以及病毒-寄主之间的互作关系，我们构建了T7和35S启动子控制下的全长cDNA克隆，但其侵染性尚需进一步证明。
Many aroid plants (family Araceae) are traditionally used for medicinal purposes in China. For thousands of years, the plants were simply collected from the wild but recently the requirements for such medicinal materials have greatly increased. Virus diseases have proved to be a particularly severe problem in vegetative cultivation of these plants.
Recently, we found Pinellia ternata (Thunb.) Ten.ex Breitenb., in Hangzhou city severely infected with a member of the genus Potyvirus. Sequencing of this virus indicated that it was most closely related to Soybean mosaic virus (SMV). We have proposed that this virus should be regarded as an aroid strain of SMV, and designated SMV-P.
To characterize the potyviruses infecting aroid plants for medicinal use in China at the molecular level, RT-PCR was done using degenerate primers specific for members of the Potyviridae to detect and identify potyviruses from several aroid plant species growing at different sites in China. Of the 20 samples of aroid plants from different sites, fragments of three distinct potyviruses were amplified from 7 samples. Of these seven samples, five were infected with SMV-P, one from Sheyang was infected with a mixture of SMV-P and Dasheen mosaic virus (DsMV), and the other one, Typhonium flagelliforme from Nanning, Guangxi, was infected with Konjak mosaic virus (KoMV-T), which was the first report of this species infecting ariod plants in China. The complete nucleotide sequence of a new Hangzhou isolate of SMV-P (SMV-PHZ3) was determined. Comparisons for each of the proteins encoded showed that, in most parts of the genome, SMV-PHZ3 had about 90% aa identity to an earlier SMV-P isolate (SMV-PHZ1) and a soybean isolate (SMV-GHZ), but that the P1 protein region of SMV-PHZ3 was substantially more similar to DsMV than to SMV soybean isolates. Phylogenetic analysis was done for the nucleotide sequence of the 3'-terminal region using the SMV-P isolates and other related potyviruses in the Bean common mosaic virus (BCMV) subgroup. The SMV-P, Watermelon mosaic virus (WMV) and SMV soybean isolates formed three distinct clusters with 100% bootstrap support but the distances between them were similar to those between different strains of BCMV and DsMV.
The 3' region (1743 nts) of DsMV-SY1 was sequenced. In phylogenetic analyses, DsMV-SY1, Vanilla mosaic virus (VanMV) and other DsMV sequences grouped together and the differences between them were quite similar to those between different strains of SMV. Serological assays revealed a weak serological relationship between SMV-P and DsMV that can be detected using antiserum prepared to the overexpressed coat proteins.
The complete sequence of KoMV-T was determined. Comparison between the amino acid sequences revealed that the P3 gene of KoMV-T was 6 amino acid longer than that of KoMV-F (the konjak isolate), which may be related to the viral pathogenicity. In phylogenetic analyses, the isolates of KoMV grouped with Zantedeschia mosaic virus (ZaMV) and Japanese hornwort mosaic virus (JHMV), although JHMV and ZaMV isolates formed separate subgroups, showing that the evolution of isolates correlated with the host and geographical distribution.
No detectable interaction was found between P1 and other viral proteins of SMV-P in yeast two-hybrid tests. Using SMV-P P1 protein as bait, an interacting plant cDNA clone was identified. Sequence analysis revealed that the insertion encoded part of a protein closely related to the cytochrome b6/f complex Riseke Fe/S gene of plants. Using 5'-RACE, the complete sequence of the Riseke Fe/S gene was obtained. The P1-Fe/S interaction was also detected by in vitro co-immunoprecipitation assay. It was confirmed by deletion mutation that the N-terminal part (aa 1-82) of SMV-P P1 interacted with the Rieske Fe/S protein of P. ternata, and that the Rieske Fe/S protein transit peptide was also involved in the interaction. Interactions between DsMV P1 and the Rieske Fe/S protein of its host Zantedeschia aethiopica, and of SMV P1 with that of soybean were also found. Although a P1-GFP fusion protein was expressed in the cytoplasm, and SMV-P interacted with the Rieske Fe/S protein of Arabidopsis thaliana, there was no obvious symptom on transgenic A. thaliana plants. It is possible that the development of symptoms is a complex process involving many viral and host proteins. In sections of virus-infected leaves, there was consistent labelling at the regions of cell cytoplasm and chloroplast using antiserum raised to the GST-PI protein while only a few (or no) scattered gold particles were seen in a control using healthy leaves.
Full-length clones of SMV-P under the control of T7 or 35S promoters were constructed. Their infectivity needs to be tested prior to their use for studying gene functions and the interactions between SMV-P and its host.
为了进一步鉴定侵染天南星科药用植物的Potyvirus属成员，我们首先用RT-PCR的方法对来自全国不同地区样品进行了检测。在供检的20个样品中，我们从7个样品中检测到了3种Potyvirus属病毒，其中5个半夏病样中含有SMV-P，1个来自江苏射阳的半夏病样含有SMV-P和芋花叶病毒(Dasheen mosaic virus，DsMV)，该分离物暂命名芋花叶病毒射阳分离物(DsMV-SY1)，另一个来自广西南宁水半夏(Tyonium flagellifrme)病样含有魔芋花叶病毒(Konjak mosaic virus，KoMV)，该分离物暂命名为魔芋花叶病毒水半夏分离物(KoMV-T)。这是KoMV侵染我国天南星科植物的首次报道。
我们测定了大豆花叶病毒杭州半夏分离物3(SMV-PHZ3)单链正义RNA基因组全序列，其全基因组由9734个核苷酸组成(不包括3’-polyA)，编码一个山3105个氨基酸组成的多聚蛋白，分子量为353.8kDa。在基因组编码的大部分蛋白中，SMV-PHZ3与大豆花叶病毒杭州半夏分离物1(SMV-PHZ1)和大豆花叶病毒杭州大豆分离物(SMV-GHZ)氨基酸序列相似性约为90％，而在P1蛋白部分，SMV-PHZ3和DsMV的氨基酸序列相似性最高。结果进一步证实，SMV-PHZ3和SMV应该是同种病毒不同分离物。同时还测定了其余5种SMV-P分离物3’术端序列，并在此基础上研究了病毒不同分离物的分子差异和进化关系。
我们测定了侵染射阳半夏的DsMV-SY1 3’末端1743个核苷酸。系统进化树进行分析表明DsMV-SY1，香果兰花叶病毒(Vanilla mosaic viurs，VanMV)和其它DsMV亲缘关系较高，成簇明显，而且DsMV分离物之间的关系和SMV分离物之间的关系很相似。Western blot分析表明SMV-P和DsMV有较弱的血清学反应。
我们测定了KoMV-T基因组全序列，其全基因组由9596个核昔酸组成，3’-末端具有poly(A)尾。KoMV-T和另一分离物F的P3蛋白氨基酸序列比较发现，两个分离物P3基因其余部分相似性比较高，但是在基因的N端度了中间部分KoMV-T比KoMV-F P3多了6个氨基酸，这可能与致病性有关。CP蛋白氨基酸序列系统进化树分析显示，KoMV分离物以及马蹄莲花叶病毒(Zantedeschia mosaic virus，ZaMV)分离物和JHMV分离物形成远源相关，与其它病毒无明显进化相关性。
用GST-P1抗血清免疫金标记发现SMV-P P1蛋白定位在细胞质和叶绿体，而在线粒体、细胞核、液泡、细胞壁等均没有观察到金颗粒标记，在阴性对照中几乎没有观察到金颗粒。P1蛋白进入叶绿体可能也是SMVP致病的一种方式。我们用酵母双杂交的方法验证P1蛋白与SMV-P编码的蛋白不存在互作。进而以SMV-P P1蛋白为诱饵从半夏酵母双杂交cDNA文库中筛选到一个候选寄主蛋白。通过质粒回收后，序列分析表明插入的部分基因与光合作用电子传递链细胞色素b_6／f复合体中Rieske Fe／S蛋白氨基酸同源性最高。用5’-RACE的方法获得该基因的全长序列。体外免疫共沉淀进一步证明P1和Rieske Fe／S蛋白互作。通过缺失突变分析表明P1-Fe／S互作的区域主要集中在SMV-P P1 N端，包括1—82位氨基酸，Rieske Fe／S蛋白的缺失突变表明Rieske Fe／S蛋白的信号肽也参与了与P1 N端34—82位氨基酸部分的互作。在大豆和马蹄莲中也同样存在SMV-P1-SFe／S和DsMV-P1-CFe／S的相互作用。共聚焦扫描显微镜下观察到GFP-P1融合蛋白在转基因拟南芥的细胞质中表达，但P1转基因拟南芥没有产生明显症状。
为了进一步研究SMV-P各基因功能，以及病毒-寄主之间的互作关系，我们构建了T7和35S启动子控制下的全长cDNA克隆，但其侵染性尚需进一步证明。
Many aroid plants (family Araceae) are traditionally used for medicinal purposes in China. For thousands of years, the plants were simply collected from the wild but recently the requirements for such medicinal materials have greatly increased. Virus diseases have proved to be a particularly severe problem in vegetative cultivation of these plants.
Recently, we found Pinellia ternata (Thunb.) Ten.ex Breitenb., in Hangzhou city severely infected with a member of the genus Potyvirus. Sequencing of this virus indicated that it was most closely related to Soybean mosaic virus (SMV). We have proposed that this virus should be regarded as an aroid strain of SMV, and designated SMV-P.
To characterize the potyviruses infecting aroid plants for medicinal use in China at the molecular level, RT-PCR was done using degenerate primers specific for members of the Potyviridae to detect and identify potyviruses from several aroid plant species growing at different sites in China. Of the 20 samples of aroid plants from different sites, fragments of three distinct potyviruses were amplified from 7 samples. Of these seven samples, five were infected with SMV-P, one from Sheyang was infected with a mixture of SMV-P and Dasheen mosaic virus (DsMV), and the other one, Typhonium flagelliforme from Nanning, Guangxi, was infected with Konjak mosaic virus (KoMV-T), which was the first report of this species infecting ariod plants in China. The complete nucleotide sequence of a new Hangzhou isolate of SMV-P (SMV-PHZ3) was determined. Comparisons for each of the proteins encoded showed that, in most parts of the genome, SMV-PHZ3 had about 90% aa identity to an earlier SMV-P isolate (SMV-PHZ1) and a soybean isolate (SMV-GHZ), but that the P1 protein region of SMV-PHZ3 was substantially more similar to DsMV than to SMV soybean isolates. Phylogenetic analysis was done for the nucleotide sequence of the 3'-terminal region using the SMV-P isolates and other related potyviruses in the Bean common mosaic virus (BCMV) subgroup. The SMV-P, Watermelon mosaic virus (WMV) and SMV soybean isolates formed three distinct clusters with 100% bootstrap support but the distances between them were similar to those between different strains of BCMV and DsMV.
The 3' region (1743 nts) of DsMV-SY1 was sequenced. In phylogenetic analyses, DsMV-SY1, Vanilla mosaic virus (VanMV) and other DsMV sequences grouped together and the differences between them were quite similar to those between different strains of SMV. Serological assays revealed a weak serological relationship between SMV-P and DsMV that can be detected using antiserum prepared to the overexpressed coat proteins.
The complete sequence of KoMV-T was determined. Comparison between the amino acid sequences revealed that the P3 gene of KoMV-T was 6 amino acid longer than that of KoMV-F (the konjak isolate), which may be related to the viral pathogenicity. In phylogenetic analyses, the isolates of KoMV grouped with Zantedeschia mosaic virus (ZaMV) and Japanese hornwort mosaic virus (JHMV), although JHMV and ZaMV isolates formed separate subgroups, showing that the evolution of isolates correlated with the host and geographical distribution.
No detectable interaction was found between P1 and other viral proteins of SMV-P in yeast two-hybrid tests. Using SMV-P P1 protein as bait, an interacting plant cDNA clone was identified. Sequence analysis revealed that the insertion encoded part of a protein closely related to the cytochrome b6/f complex Riseke Fe/S gene of plants. Using 5'-RACE, the complete sequence of the Riseke Fe/S gene was obtained. The P1-Fe/S interaction was also detected by in vitro co-immunoprecipitation assay. It was confirmed by deletion mutation that the N-terminal part (aa 1-82) of SMV-P P1 interacted with the Rieske Fe/S protein of P. ternata, and that the Rieske Fe/S protein transit peptide was also involved in the interaction. Interactions between DsMV P1 and the Rieske Fe/S protein of its host Zantedeschia aethiopica, and of SMV P1 with that of soybean were also found. Although a P1-GFP fusion protein was expressed in the cytoplasm, and SMV-P interacted with the Rieske Fe/S protein of Arabidopsis thaliana, there was no obvious symptom on transgenic A. thaliana plants. It is possible that the development of symptoms is a complex process involving many viral and host proteins. In sections of virus-infected leaves, there was consistent labelling at the regions of cell cytoplasm and chloroplast using antiserum raised to the GST-PI protein while only a few (or no) scattered gold particles were seen in a control using healthy leaves.
Full-length clones of SMV-P under the control of T7 or 35S promoters were constructed. Their infectivity needs to be tested prior to their use for studying gene functions and the interactions between SMV-P and its host.
引文
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